heteropteranany member of the insect order Heteroptera, which comprises the so-called true bugs. (Some authorities use the name Hemiptera; others consider both the heteropterans and the homopterans to be suborders of the Hemiptera.) This large group of insects, consisting of more than 40,000 species, can be recognized by an X-shaped design on the back, which is formed by the wings at rest. A combination of features—sucking mouthparts adapted to pierce plant or animal tissues and a hardened gula (underside of the head)—separate the heteropterans from all other insect orders. Although most species of Heteroptera are terrestrial, a few are aquatic. Some species, which feed on plant juices, are serious pests of cultivated crops; other species are predacious and benefit man by destroying various pests. There also are heteropterans that act as carriers of disease.

Heteropterans can be divided into three large groups on the basis of general habitat: the water-dwelling Hydrocorisae (water boatmen, back swimmers, water scorpions, giant water bugs, and creeping water bugs); the surface-swimming and shore-dwelling Amphibicorisae (water striders, marsh and water treaders, shore bugs, and velvet water bugs); and the Geocorisae, a large group of land bugs (plant bugs, bedbugs, assassin bugs, anthocorid bugs, lace bugs, ambush bugs, stinkbugs, burrower bugs, stilt bugs, and fire bugs).

General features

Size range and diversity

Like any other biologically successful group of organisms, the heteropterans are prolific and diverse and have adapted to a variety of habitats. They range in length from more than 100 mm (3.9 inches) to less than 1 mm (0.039 inch) and have invaded habitats from dry land to water. One of the few insect groups with aquatic adults capable of obtaining adequate oxygen from the water, heteropterans include the only insect species that spend their lives on oceans far from land. Some heteropterans are nourished by blood of animals ranging from mites to man; others feed from the sap of plants as diverse as giant redwoods and algae. Some suck surface fluids (e.g., nectar), some pierce tissues to suck sap or blood, and others obtain nourishment from dried seeds. Many forms live on open surfaces and escape enemies by running, flying, or remaining inconspicuously motionless. Some seek food and shelter in natural crevices; others actively burrow into the soil or seek out the nests of animals.

Distribution and abundance

Heteropterans are most abundant in the tropics, decreasing in both individuals and species to limits northward beyond the Arctic Circle and southward almost to the Antarctic Circle. Every major land mass harbours different species, and migrations to new habitats may be aided by natural agents (e.g., wind, birds, floating debris) or by man. Although heteropterans have been carried throughout the world, only a few species have become established in many lands. Unique among insects are some water striders (Gerridae), which are at home on the open ocean between approximately 40° north latitude and 40° south latitude and may not approach land for several generations.

More than 40,000 species of heteropterans are known. Most families containing 150 or more species are represented in every zoogeographic region. Two exceptions are the Phymatidae (ambush bugs), with no species known in Australia or the Pacific Islands, and the Plataspidae, with no New World species. Families with the fewest species have the most restricted ranges: Aphylidae, Hyocephalidae, and Lestoniidae are restricted to Australia and contain among them only five species. The Velocipedidae of the Orient and the Vianaididae of the American tropics number four species each. In general, zoogeographic restriction is most evident at the subfamily, tribal, and generic levels.

Importance

Heteropterans have complex and important roles in the balance of nature. The majority of them occupy an intermediate position in the ecological food chain; they use food producers (plants) and serve as food sources for parasites and other animals. A few species utilize plant-feeding insects as food.

Beneficial aspects

Heteropterans often serve as food for man and other animals. In some Latin American countries eggs of certain aquatic bugs are collected by providing submerged mats of hollow reeds or straws as egg-laying sites; the eggs are then dried and made into cakes. Chickens, turkeys, hogs, and other domestic stock consume available heteropterans. Aquatic bugs are an important food source for fish. Wild birds and mammals, from the wrens and shrews to the turkey and grizzly bear, utilize available heteropterans in their diets.

Species that attack unwanted plants and feed on the eggs, immature stages, and adults of injurious insects include most assassin bugs, nabid bugs, anthocorid bugs, stinkbugs, and plant bugs. These heteropterans are used in biological control of noxious weeds and injurious insects.

Harmful aspects

In contrast, some heteropterans (e.g., plant bugs, chinch bug, lace bugs, stinkbugs) damage both wild and domestic plants by sucking sap or injecting tissue-killing fluids. Insertion of a beak or eggs into plant tissue harms the protective layers and allows bacteria and other disease-causing organisms to infect the plant. Although heteropterans are not the major agents for transmission of plant viral diseases, a few species (Piesmatidae) can carry viral diseases of sugar beets and related crops in North America and Europe.

Heteropterans can affect man in several ways. They are common household pests and can spoil the taste of some fruits (e.g., raspberries contaminated by stinkbugs). But more importantly, some can attack man directly and inflict painful bites as well as introduce disease-causing organisms. The injection of saliva or poison may cause allergic reactions in susceptible persons. Sometimes when they cannot find plants, phytophagous insects probe moist surfaces (e.g., perspiring skin) in search of appropriate food fluids. Transmission of trypanosomes, which cause Chagas’ disease in the American tropics, occurs through cone nose bugs (Reduviidae), so-called because of the shape of their head. The insect receives trypanosomes when it feeds on the blood of an infected human. The trypanosome passes part of its life cycle in the insect and again becomes infective to man. Instead of injecting them into a new victim, however, the insect deposits trypanosomes (in its excrement) on the skin of a potential victim. The trypanosomes can enter the bloodstream only through mucous membranes or breaks in the skin (e.g., those that may result from scratching the site of the bite).

Natural history

Life cycle

General features

In tropical regions, where heteropterans are most abundant, populations are influenced by factors such as rainfall, host plant development, and internal rhythms. In addition to these, heteropterans in temperate regions are influenced by seasonal temperature fluctuations.

Heteropterans generally produce one or two generations a year in temperate regions. The dormant stage, which varies from species to species, is dependent in part on the food source of the species. For example, a heteropteran that feeds on perennial plants often lays eggs that remain dormant through the winter in or on the host plant. The nymphs, which hatch in the spring, then have appropriate leaves and succulent stems to furnish the sap necessary for growth. In contrast, heteropterans that feed on annual plants have a dormant winter adult stage that flies to a new host plant each spring after seeds germinate. Species that have short nymphal and adult stages, which coincide with certain plant characteristics (e.g., short-lived flowers), may pass 10 or 11 months as eggs. Among predatory heteropterans, those whose prey is found on plants or in flowers winter as adults; those whose prey is found on the ground may be dormant as nymphs or adults. Aquatic heteropterans may spend the winter as adults, nymphs, or both, whereas semiaquatic forms are dormant as adults.

Metamorphosis

Heteropterans undergo gradual metamorphosis (hemimetabola). Primary feeding and energy storage, as well as development of adult structures, take place in the nymphal stages. The adults seek mates and potential food sources and are responsible for initiating future generations. The developing embryo, incapable of obtaining food or defending itself, is encapsulated within a protective egg shell that contains adequate food. The nymph, which lacks the locomotor and reproductive ability of the adult, also lacks the hard and rigid extensive exoskeleton necessary for strong muscle action. As a result, however, its body can distend to accommodate large quantities of food. In addition, loss of moisture through the thinner, more membranous body cover is offset by the high water content of the nymph’s food.

Reproduction

Eggs

Reproduction occurs when eggs are fertilized as they pass through the female ducts by spermatozoa stored in special receptacles in the female. A notable exception to this method occurs in the bat bugs (Polyctenidae). In this case shell-less, yolkless eggs are fertilized in the female ovariole where they remain, while the developing embryo is nourished by special maternal cells through a placenta-like structure consisting of tissues from both parent and offspring. The bat bug nymph leaves the mother’s body only after reaching an advanced stage of development. Heteropteran eggs generally are moderate to large in size relative to adults and have a smooth to finely sculptured surface, sometimes with a colour pattern and often with slender projections. Eggs contain sufficient nutrients to permit the embryo to develop into a free-living, sexually immature, wingless nymph. Just before an egg hatches, the shell becomes translucent and reveals the segmentation and strongly coloured structures (e.g., eyes) of the nymph inside.

Eggs may be laid singly or in clusters. Eggs of plant-eating species generally are glued to a surface or inserted into the tissues of a selected host plant. Eggs of predatory forms often are laid near prey. Aquatic bugs may lay their eggs above or below the water’s surface. Some bugs have special habits; for example, eggs may be glued in a large cluster to the back of a heteropteran male (Belostomatidae) or to the body of a crayfish as in Corixidae.

Nymphs

A newly developed nymph uses one or both of the following mechanisms to escape the egg: a cuticular spine on the head, sometimes known as an egg burster; or internal hydrostatic pressure created by forcing fluids (sometimes in the head) against the site of egg rupture. The pattern of rupture may be controlled by a line of weakness in the egg; in some cases a flaplike operculum lifts back to allow the nymph to escape. After withdrawing itself from the egg, the newly hatched nymph seeks the food source essential for its development. Some nymphs leave the egg with a sufficient store of nutrients to allow them to pass through their first molt (or ecdysis). This is especially important to predatory species whose newly hatched nymphs might have to spend considerable time searching for prey.

The wingless nymph must rely on mimicry or disguise to prevent detection by enemies. Relatively inactive nymphs are often the same colours or shapes of objects in their immediate environment. Other types of nymphs disguise their bodies with a layer of small particles collected on a coat of sticky body hairs or exhibit contrasting colours that break up the body outline. Some active nymphs confuse predators by alternating short quick dashes with periods of no motion or deceive predators by simulating the body shape, colour, or erratic gait of ants, with which nymphs sometimes are associated (see coloration; mimicry).

Lacking a pupal stage in which to change into an adult, the flightless nymph must sustain itself while it gradually transforms into an adult. As the nymph feeds and grows, it discards through five successive molts (ecdyses) the growth-restricting outer layers of its exoskeleton. Wings appear first as slight outpocketings at posterior regions of the mesothorax and metathorax during the third instar (interval between molts) and enlarge during the fourth and fifth instars. However, the wings are not fully functional until the last ecdysis, when the nymph becomes an adult. At the time of the last molt, other changes occur: the head and thorax assume new shapes; the number of segments in antennae and tarsi may change; scent glands on the top of the abdomen cease functioning and are replaced by metathoracic scent glands; and the external genitalia and the internal reproductive organs become functional.

Adults

Flight is common among adult heteropterans. Although they are capable fliers, no species has developed the aerial efficiency of some other insect groups (e.g., dragonflies, flies). Predatory heteropterans lie in wait for a potential victim to wander close enough to be captured, instead of overtaking their prey in flight. Winged adults may rely on flight or inaction and the optical deceptions of colour arrangement and body form utilized by the nymphs to escape enemies. Flight also enables the two sexes to come together and provides an effective means for seeking favourable egg-laying sites.

Behaviour

Sound production and reception

Due to their small size, heteropterans cannot produce the conspicuous sounds typical of katydids, crickets, and cicadas. Sound production by specialized body parts (also called sonification or stridulation) is common among heteropterans but seldom loud enough to attract human attention. Heteropterans produce sounds by moving one roughened member over a roughened area of body surface. Sometimes both roughened parts consist of a series of minute, closely-set, parallel furrows or ridges called a strigose area, strigil, file, or rasp. Sometimes the second part of the mechanism consists of a series (called a plectrum) of minute pegs, setigerous tubercles, or an upturned edge of a sclerite (hard body plate). The plectrum may be on the movable body member or on the fixed body part. One or both sexes, and sometimes even the nymphs, of a species may stridulate. Some sounds appear to have courtship significance. The presence and form of sound-producing mechanisms are useful in the classification of Heteroptera. Although many have similar sound-producing structures, the whirring or buzzing noise common to heteropteran flight is caused by the rapidly moving flight mechanism.

Mechanisms for detecting airborne and possibly waterborne vibrations (or sounds) are known in Heteroptera: auditory organs known as Johnston’s organs are found in the antennae of all species, and the tympanal (stretched membrane) organs are known in several families of aquatic bugs. A variety of receptors, especially for detecting substrate vibrations, have been described for members of this order.

Feeding

Heteropterans are essentially nonsocial insects. Newly hatched nymphs occasionally remain together for a short time, clinging to the shells of the eggs from which they have hatched; for example, plant feeding (phytophagous) nymphs may remain together because they hatch on their food source and need not search for it. In contrast, however, most predatory heteropterans encounter their prey by chance, and, in this case, dispersion increases individual opportunities for finding prey and for escaping other predatory heteropterans. Apparent coordinated migrations reported for chinch bugs generally result from a sudden disappearance or failure of a food source.

Ecology

Heteropteran behaviour, governed by natural instincts, includes patterns that enable individual species to live within specific environments. The patterns are so similar for all members of a family that each family can be characterized as essentially phytophagous or predatory, and terrestrial, semiaquatic, or aquatic. Most heteropterans rely on atmospheric oxygen for respiration.

Habitats

Aquatic Heteroptera vary in their independence of atmospheric oxygen. Members of one of the nonswimming families (Nepidae) submerge just below the water’s surface and obtain oxygen through a slender, terminal breathing tube that breaks the surface film. Some swimming families (Corixidae, Notonectidae) dive deeply but must surface periodically to replenish air supplies either stored in chambers formed against body surfaces by the folded wings or entrapped by rows of dense hairs folded against the body. These insects swim to overcome the buoyancy of the renewed air supply, grasping underwater objects to anchor themselves. Heteropterans with air storage chambers are independent of oxygen dissolved in the water; therefore, they survive in warm springs or polluted waters almost devoid of oxygen. Another family (Naucoridae) contains species that spend their entire lives below the water’s surface and can obtain oxygen from that dissolved in the surrounding water. Water insulates its inhabitants from cold temperature extremes, allowing aquatic species to exhibit considerable activity throughout the year. Chilled water holds more dissolved oxygen than warm water, and air is available also between the water’s surface and the ice layer during winter months in temperate regions.

Semiaquatic bugs literally walk on water in that they are supported solely by the water’s surface film. Their small, slender bodies have little bulk to support. Long, thin middle and hind legs spread the points of contact with the water, with the tarsus of each leg causing only a slight dent in the surface film. The shorter front legs may rest on the water or capture and hold prey. Wetting of the body during submersion is prevented by a thin layer of air trapped by the fine, dense, water-repellent (hydrofuge) pile that covers the insect. Many surface dwellers live in quiet water, whereas others prefer flowing water, even rapids, where considerable strength is needed to overcome the current. In some cases, a bug may anchor the legs on one side of its body to some fixed object in a stream and wait for food carried to it by the rushing water. Adults survive the winter beneath objects on land either close to or some distance removed from a body of water.

Although shore bugs normally frequent shore lines, they occasionally make short forays onto the surface of adjacent bodies of water. Shore bugs generally are heavier bodied than surface dwellers. Their strong, and often long, legs enable them to run quickly across ground, rocks, and other exposed surfaces; short rapid flights are not uncommon. All shore bug families are predatory. Their front legs often are thickened distinctly and sometimes spined for grasping prey. Adults generally overwinter beneath convenient objects.

Terrestrial heteropteran species are more abundant than either aquatic or semiaquatic forms and have a variety of habits and life histories. Some species wander over leaves, flowers, and other surfaces exposed to sunlight, whereas others prefer protected areas such as crevices and the underside of foliage, rocks, logs, and loose bark. Some species, which are active during the night, hide only during the day, whereas other groups (e.g., Aradidae) live in obscure habitats and leave only if the environmental conditions become unsatisfactory. Members of one family (Cydnidae) burrow by cutting soil in front of them and shoving it behind and live in moving underground chambers. Species in at least two families (Nabidae, Reduviidae) live in spider webs and skillfully avoid entanglement as they eat trapped insects or, in some cases, the spider itself.

Relationships of heteropterans to plants vary. Plants may serve simply as a site on which to stand or move, as a lure for plant feeding insects on which predatory bugs feed, or, most importantly, as a source of nourishing sap for phytophagous heteropterans. A few species can cause modifications in plant growth. For example, a feeding insect may cause leaves to roll up around it. In Europe certain lace bugs (Tingidae) insert their eggs into flower buds, which enlarge and house developing lace bug nymphs. A new adult lace bug does not need jaws to gnaw its way out because it simply waits for the bud to begin to open.

Food habits

Predation is common throughout the Heteroptera, sometimes as a family-wide feeding pattern, sometimes as a deviation within an otherwise phytophagous family. In the suborder Hydrocorisae all families except one (Corixidae) are predatory on small aquatic animals, including small fish, tadpoles, and frogs. The exception has been known to add mosquito larvae to its usual algal diet. The suborder Amphibicorisae, without reported exception, contains forms that prey on plankton, small arthropods, or worms. In the suborder Geocorisae, many families of the series Cimicomorpha (e.g., Reduviidae, Nabidae, Phymatidae) are predatory, generally on other insects. However, a few assassin bugs (Reduviidae), bedbugs (Cimicidae), and bat bugs (Polyctenidae) seek the warm blood of birds and mammals. In addition, some members of plant feeding families (including the Miridae) attack other arthropods; a few families (Pentatomidae, Lygaeidae) of the series Pentatomomorpha, fundamentally plant feeders, contain insect-eating species. The original food habit of this order is not yet clear, but even in some avidly predatory species, the first and second nymphal instars may be plant feeders. In capturing its prey, a heteropteran may use front legs modified for grasping or insert feeding stylets. In either case the victim is paralyzed quickly by an injected salivary secretion.

Any insectan stage may fall to the hungry predator. Insect eggs are a favourite food for many small (Anthocoridae) or very young heteropteran predators. Caterpillars and other larvae commonly are attacked by predatory Heteroptera, and it is estimated that one stinkbug (Pentatomidae) may destroy more than 200 caterpillars during its lifetime. One stinkbug, long known to feed upon bedbugs, was recommended for use in control as early as 1776.

Other arthropods also may be attacked. One anthocorid bug (Anthocoridae) may destroy an average of 33 mites per day. In some instances size of prey is no barrier. In Sri Lanka a small assassin bug (Reduviidae) only 4 mm (0.157 inch) long can paralyze a giant millipede 150 mm (5.9 inches) long.

Associations

Individual heteropterans may associate accidently on a food source. On quiet pools of water, water striders (Gerridae, Veliidae) often assemble in milling “schools” of a few to hundreds of individuals that quickly reassemble if dispersed. Sometimes a female will reassemble disturbed batches of eggs. In cases where the female lays her eggs on the back of a male, the parental responsibility is his.

Members of one family (Termitaphididae), unable to live alone, share the nests of certain termites and feed on the fluids of the fungi growing there. In exchange, a substance secreted from pores on the backs of these bugs is eaten by the termites. Reports of heteropteran associations are difficult to assess and may reflect coincidental use of the same environment.

Form and function

General features

Heteropterans have features typical of all insects and special adaptations that differentiate them from other insect groups. The head tests new environments through light perceptions by compound eyes and simple eyes (ocelli) and through both scent and sound perceptions by the antennae. The mouthparts are located conveniently for quick exploitation of newly discovered food sources. The thorax bears support and transportation mechanisms, legs, and wings. An important function for the third body region, the abdomen, is reproduction. Sutures divide the body surface into sclerites (hard plates), which may be fused or separated by flexible membranes. Hairs, scales, or a variety of glandular secretions may adorn at least some areas of the body. The body surface may be raised into spines of a variety of shapes, knobs, or tubercles; tubercles often are setigerous; i.e., glandular with a straight or curved filament or bristle at the tip. Some of these projecting structures, which have bases associated with specialized cells, probably have a sensory function and respond to vibrations of air, substrate, or touch. Certain long hairs, each arising from a ringed socket, are called trichobothria, and their presence and arrangement on various parts of the body have taxonomic significance.

Distinguishing features

The covering of the head is a sclerotized ring tapering from the flexible neck forward or downward and ending in three lobes: a jugum on each side and a central clypeus (or tylus). The area above and between the eyes (called the vertex) may bear a transverse pair of simple eyes (a median one is never present in Heteroptera). The hard lower surface of the head is called the gula (hypostomal bridge).

Mouthparts

The piercing–sucking mouthparts are composed of a troughlike, four-segmented labium in which lie four stylets; these are modified mandibles and maxillae. Each of the hairlike maxillae has two major grooves plus minor grooves and ridges along its median surface. When brought together and locked by the minor grooves and ridges, the two major grooves form the left and right halves of two separate tubular canals that extend the length of the maxillae. The anterior canal is the food tube through which fluids are drawn by a sucking pump (described below); the posterior tube is the salivary canal through which digestive and certain other fluids may pass into a food source. These fluids stun or kill the prey, prevent coagulation of blood, and initiate the process of digestion. Along the sides of the maxillary stylets are slender mandibles with serrated tips. The mandibles and maxillae alternately advance into the food tissue until the appropriate fluid is reached by the maxillae, which is when ingestion begins. Since the labium does not penetrate the food tissue, the stylets must extend beyond the protection of the labial groove. This extension may be accomplished in one of two ways. The labium may be shortened by jackknifing some of its segments, leaving only the apical and sometimes the basal segments to guide the exposed stylets. In some species the extensible stylets are much longer than the labium. When not extended, the stylets either are coiled or looped in a chamber located in the labium or the head or are found in a long sheath that extends backward into the heteropteran’s body cavity.

The sucking pump is a chamber whose front wall is folded lengthwise into the mouth cavity. Contraction of certain muscles increases the size of the cavity, decreases the pressure in it, and allows the liquids to flow through the food canal into the pump. When the muscles relax, the size of the chamber decreases gradually and pushes the food toward the true mouth, which opens into the esophagus.

Thoracic features

Each of the three somites of the thorax bears a pair of legs, and both the meso- and metathorax bear a pair of wings. The large conspicuous part of the thorax between the head and the wings is called the pronotum (i.e., the expanded top part of the prothorax). Projecting from beneath the hind margin of the pronotum is the triangular, sometimes U-shaped, scutellum. Along the side margins of the scutellum may be a groove or fold (the frenum). In repose, the inner edges of the front wings hook onto the frenum. On each side of the metathorax may be a pore (ostiole) that provides an outlet for a highly volatile, repellent fluid secreted by internal scent glands. The ostiole may be surrounded by an elevated ridge (peritreme), and then by an extensive, roughened, evaporative area.

Legs

The legs may be modified in various ways to serve specific functions. Legs may be lengthened or expanded to aid in camouflage. Sometimes the hind femurs and their accompanying leg muscles are adapted for jumping. The middle and hind legs may be flattened for swimming, or their effective width increased by a marginal fringe of long hairs that spread during a power stroke and flatten against the leg on the recovery stroke. The front legs, when specialized for grasping prey, have tibia and tarsus adapted to fold tightly against a thickened femur. Stout hairs or spines along one or both of the opposing surfaces of this grasping leg prevent a victim from slipping free. The males of many species have hind legs that are thickened or spined to hold females during mating. Lacking mandibulate mouthparts with which to clean their antennae, many Heteroptera have a comblike series of hairs on the end of the front tibia, across which the antennae may be dragged. Certain genera (Nabidae, Reduviidae) that live suspended slothlike from the underside of spider webs have long, foldable claws.

Wings

The texture of each of the two pairs of wings is distinct in heteropterans. The mesothoracic or forewings (called hemelytra, elytra, or tegmina) are stiff and have an oblique line that abruptly separates the leathery basal half from the membranous apical half, while the metathoracic or hindwings are thin, delicate membranes. At rest the forewings are folded over the hindwings. The membranous tips of the forewings overlap, so that the fragile hindwings lie beneath them. In macropterous individuals with wings of normal size, the wings approach or extend beyond the apex of the abdomen. Some species have wings that are brachypterous (slightly to moderately reduced), micropterous (very small), or absent. Both pairs of wings are absent in one family (Termitaphididae), apparently an adaptation to the termite tunnels in which they live. In the bat parasite family (Polyctenidae) the budlike character of the nymphal wings is carried into the adult stage with no further development, and these insects are considered to be wingless. The front pair of wings of the bedbug family (Cimicidae) is reduced to short, nonfunctioning pads, and the hind pair is lost completely. Although these heteropteran families are the only examples of a general pattern of wing loss or reduction, some individual species show a variety of wing developments (alary polymorphism) or aptery (winglessness) in one or both sexes. Species with sexual dimorphism in wing length have females with reduced wings or none at all; apparently the retention of flight in the male assures his ability to move to the female.

Heteropteran wings, especially the front pair, may be modified for specific functions. Their presence above the abdomen provides protection for the soft parts beneath. More effective protection is provided in forms with fused forewings. Some forms can couple their wings by means of a fingerlike projection (jugum) on each forewing, thus enhancing their flying ability. The colour and texture of the forewings may be modified so that an insect resembles bark. The lace bugs (Tingidae), so-called because they have a lacelike pattern of many fine, interconnecting wing veins, resemble the fine veins on the undersides of the leaves on which they feed. In some species, bright or contrasting wing colours announce unpalatability to an enemy.

Abdomen

The abdomen contains 11 segments, with the 10th and 11th, and sometimes others, fused together. Sensory appendages (cerci) are never present in this order, but the sensory function may be performed in part by several apparently tactile hairs along the underside of the abdomen. The terminal abdominal segments of the male are adapted primarily for transfer of sperm, those of the female for reception of sperm and egg laying. Females that insert eggs into plant tissue have an ovipositor under the tip of the abdomen, which consists of blades that slice plant tissue and guide the eggs into the slit. In females that glue eggs to a surface, the ovipositor usually is reduced to a noncutting egg guide.

Internal features

Digestive system

The three embryological regions of the alimentary canal become modified by constrictions, dilations, and outpocketings. The stomodaeum becomes a short esophagus that connects to the midgut. The midgut frequently differentiates into four distinct regions, two dilations connected by a short, narrow tube and a second tubular section that gives rise to slender, terminally closed, tubular outpocketings, called mycetomes. The cavities of the mycetomes are filled with bacteria, whose role may be either to supply essential nutrients or to inhibit development of other bacteria. The bacteria, hereditarily transferred from mother to offspring, migrate to the ovarian follicle of the female and invade the eggs. As soon as an embryo begins to develop a gut, the bacteria appear in it.

Respiratory system

The respiratory system consists of longitudinal tracheal trunks that branch internally and communicate with the external air through ten pairs of holes called spiracles. Respiration under water presents special problems. Young aquatic nymphs may respire exclusively through the thin body wall. Adult aquatic insects, with their hard body walls, must rely on tracheal respiration and modifications of the spiracles and external skeleton surrounding them, which serve to exclude water while admitting oxygen. In some species the spiracles are sunken and protected by a circlet of hydrofuge (water repellent) hairs or by a fine-pored membrane, whereas in other aquatic heteropterans the spiracles are closed and only one or two pairs function in association with either diving air storage chambers or the base of a snorkel-like breathing tube that breaks the surface film of the water. Although air storage chambers absorb some dissolved oxygen from the surrounding water, they gradually diminish in size as the nitrogen contained in them dissolves outward into the surrounding water. Unless they are replaced by frequent trips to the surface, air supplies may be lost completely, thus allowing water to reach and enter the spiracle, which would result in drowning. A few forms (Naucoridae) have “plastron” respiration. The plastron is actually a modified air storage chamber that consists of air in a series of cuticular grooves radiating from a spiracle. This air is continuous with tracheal air and is held in each groove by hydrofuge hairs, so that its volume does not change. Sustained absorption of dissolved oxygen by the plastron eliminates the need for contact with atmospheric air as long as the surrounding water is well oxygenated.

Other systems

The skeletal, muscular, circulatory, and excretory systems are structurally and functionally typical of a winged insect with the exception of some adaptive variations. The piercing-sucking mouthparts require protractor and retractor muscles for insertion and withdrawal of each stylet, and dilator muscles to enlarge the sucking pump cavity. General blood flow throughout the body is provided by the four- or five-chambered heart, but circulation in legs and wings requires special devices (e.g., the pulsating membranes located within the base of each tibia and sometimes in the tarsus). Rhythmical waving of pulsating membranes causes a pattern of blood flow to the tip of the limb and back. The number of excretory tubes (i.e., malpighian tubules) generally is four.

The nervous system consists of three interconnected parts known as the central, visceral (sympathetic), and peripheral systems. The ventral nerve cord exhibits considerable concentration of ganglia near the head end (cephalization). In all species the abdominal ganglia have migrated into the thorax. Most members of the order have three ganglia on the ventral nerve cord; the subesophageal, the first thoracic, and a mass that results from fusion of the second and third thoracic ganglia with all the abdominal ganglia. Complete cephalization of the ventral nerve cord occurs in a few Heteroptera where all the ventral ganglia, including the subesophageal, unite into a single ganglionic centre.

The heteropteran reproductive system consists of a pair of gonads whose ducts unite to form a single tube leading to the exterior near the posterior end of the abdomen. In most species spermatozoa are transferred directly into the female reproductive tract and stored in a spermatheca (outpocketing of female tract) until fertilization occurs (as the egg is laid). In two families (Cimicidae, Anthocoridae) the female has an organ separate from the reproductive tract to receive the spermatozoa. This organ is a rounded internal pouch associated with a slit on the underside of the abdomen and is called the organ of Ribaga. During mating the spermatozoa are deposited in this pouch. They then penetrate the pouch wall, travel through the body cavity, and burrow into the spermatheca, remaining there until needed to fertilize the eggs. Excess spermatozoa are absorbed as nutrients by special cells in the female.

Evolution and paleontology

The fossil record is too poor to offer significant help in reconstructing the phylogeny of the Heteroptera. Their small size, the fragile nature of dead individuals, and a preference for habitats that seldom are conducive to fossilization have resulted in few fossils. Often only wings or parts of wings are represented. Known fossils do reveal, however, that during the Permian Period (more than 225,000,000 years ago), two major lines existed. The order Protohemiptera, which revealed a number of features common to homopterans and heteropterans, including stylet-like mouthparts assumed to be piercing-sucking, actually represents an unrelated parallel development.

The appearance of the Heteroptera as a line distinct from the Homoptera also is represented in Permian fossils. The oldest known member of the order Heteroptera (from the Permian in Australia) represents the only family (Paraknightiidae) with no living species. Fossils from the next geological period, the Triassic, reveal that the major forking of the heteropterans into aquatic and terrestrial branches had already occurred near the beginning of the Mesozoic Era. Several modern aquatic family types are recognizable in the Mesozoic. The bulk of modern terrestrial heteropterans appears in the fossil record by the Tertiary Period Paleogene and Neogene periods of the Cenozoic Era.

Classification

Distinguishing taxonomic features

Modifications of mouthparts and wings are generally accepted as important characteristics by which to divide the class Insecta into orders. One peculiar type of mouthpart, possessed by more than 50,000 insect species, is the piercing-sucking type in which the labium forms a trough containing two pairs of threadlike stylets (highly modified mandibles and maxillae). All modern insects with mouthparts of this type are closely related to each other but not to other insects, none of which possess truly comparable modifications. Some entomologists consider all insects with piercing-sucking mouthparts as members of a single order, the Hemiptera (sometimes called Rhyngota or Rhynchota), with two suborders Homoptera and Heteroptera separated traditionally by texture and resting position of the forewings and by the apparent origin of the beak. Other entomologists, while recognizing the proximity of relationship between these two groups, consider that the relationship is of superorder value and that features separating the two groups are of sufficient magnitude to warrant full ordinal status for each group.

Separation of the two groups is reflected by differences in several important functioning body parts. The gula of the Heteroptera is a hard bridge that separates the mouthparts from the prothorax. In the homopterans the gula is small and membranous or absent, allowing the base of the mouthparts to abut or fuse to the prothorax. The forewings of heteropterans are generally divided into basal and apical halves of different texture. Homopteran forewings, on the other hand, are generally of similar texture for their full length. Scent glands are present in all terrestrial and some aquatic nymphs and adults of the Heteroptera. Homopterans lack scent glands.

Reduction and loss of the gula in homopterans, coupled with the migration of the mouthparts to the prothorax, were significant changes in functional specialization. In some homopteran families of the suborder Sternorrhyncha, the mouthparts have fused to the prothorax and remain attached to it if the cranium is removed. These and other basic characters that began to differentiate in the geologically remote Permian time (when many other insect orders were evolving) indicate an important branching of the phylogenetic line of homopterans and heteropterans and substantiate their separation at the ordinal level.

The heteropterans are divided into suborders on the basis of habitat, antennal structure, ability to produce sound (stridulate), and presence and arrangement of long hairs (called trichobothria) on various parts of the body. Division into families is based on structural differences and adaptations for specialized ways of life.

Annotated classificationOrder Heteroptera (true bugs)Wings, when present, usually number four that may vary in size; basal portion of forewings thick and leathery, apical portion membranous; hindwings membranous, slightly shorter than forewings; wings at rest held flat over abdomen; piercing-sucking mouthparts arise from front part of head; compound eyes usually well developed; two simple eyes, or ocelli, present or absent; scent glands usually present; gradual metamorphosis; widely distributed, terrestrial and aquatic species number more than 40,000.Suborder Hydrocorisae (or Cryptocerata)Neither cephalic nor abdominal trichobothria; antennae 4-segmented, shorter than head, usually in grooves on underside of head; semiaquatic (Gelastocoridae, Ochteridae) or aquatic (all other families); swimming members with fringe of swimming hairs on hind legs; aquatic members lay eggs in or on submerged exposed objects, are relatively active under water during winter; probably all males, and some females, stridulate; important food for fish, some water birds.Family Corixidae (water boatmen)Head overlaps front of prothorax; scutellum small, usually concealed by pronotum; front legs short, 1-segmented tarsus with fringe of long hairs beneath for gathering food; middle legs long, slender, for submerged anchoring; males with stridular pegs on front femurs and tibiae of legs, a polished black strigil above on asymmetrical abdomen; females nonstridulatory, abdomen symmetrical; colour pattern commonly alternating light and dark transverse stripes; inhabit fresh, brackish, or salt (Great Salt Lake) waters; use diving air stores; phytophagous on algae and diatoms, but may eat insect larvae; more than 300 species; in all major faunal regions.Family Notonectidae (back swimmers)Head not overlapping prothorax; scutellum large, exposed; all legs long and slender, hind pair longest; male stridulating mechanism involving front legs and labium; inhabits fresh and mineral waters; diving air stores trapped under hairs on underside of abdomen, hence inverted position in swimming; bite painful to man; predatory on insects, small crustaceans, fish, and tadpoles; almost 200 species; in all zoogeographical regions.Family PleidaeVery small, stout, convex above; scutellum large, exposed; hindwings reduced or absent; head and pronotum partially or wholly fused; inhabit fresh waters; about 20 species; in all zoogeographic regions.Family HelotrephidaeLike Pleidae; about 15 species; only in Asia, Africa, and South America.Family Nepidae (water scorpions)Body stout to very slender; legs long, generally slender, front pair adapted for grasping; abdomen of most nymphs and adults with snorkel-like breathing tube at tip; nymphs and adults stridulate by opposable files on anterior coxae and coxal cavity plates; inhabit fresh water; swim poorly or not at all; predatory; principal food, other insects; nearly 200 species; in all zoogeographical regions.Family Belostomatidae (giant water bugs and toe biters)Large, some species exceeding 109 millimetres (4 inches) in length; broad, flattened body; middle and hind legs markedly flattened, with swimming hairs; inhabits fresh water; may lie in mud or debris on bottom; can inflict severe bites; predatory; aggressive, strong swimmers; grasping front legs capture insects, fishes, frogs; approximately 100 species; in New World, Asia, and Africa (including Madagascar).Family Naucoridae (creeping water bugs)Head large, inserted into thorax; scutellum large, exposed; front femurs enlarged, tibiae opposable; middle and hind legs slender, sometimes with fringing hairs; inhabit fresh water, often warm springs; generally use diving air storage chambers or “plastron” respiration; predatory mostly on mollusks and insects; about 150 species; most common in tropical continents, ranging north into Europe and North America; subfamily Aphelocheirinae uses plastron respiration, sometimes elevated to family rank.Family Gelastocoridae (toad bugs)Broadly oval in shape, roughened surface, elevated protuberant eyes; resemble tiny toads; beak reaching only to coxae of front legs; broadened front femurs and opposable tibiae form effective grasping organ; middle and hind legs slender for running, lack fringing hairs; inhabit shores, burrows into mud or sand to lay eggs; predatory principally on small insects; about 100 species; in all zoogeographic regions.Family Ochteridae (velvety shore bugs)Similar to Gelastocoridae in general appearance; body surface smooth, with velvety pile; beak reaches abdomen; found among plants growing along margins of ponds and streams; predatory upon other insects; about 25 species; in all zoogeographic regions.Suborder AmphibicorisaeTrichobothria on head but not on abdomen; members generally live on the water’s surface film or on mud or sand; predatory on tiny animals, including injured or freshly-killed insects; nonstridulating except for both sexes of genus Stridulovelia (Veliidae); within one species individuals may vary from winged to wingless.Family Gerridae (water striders; pond skaters)Slender bodied; legs long, slender, front pair short, used for support or grasping prey; middle pair longest, used for propulsion over water surface; hind femurs extends beyond tip of abdomen; tips of tarsi project beyond insertion of claws; body covered below with hydrofuge hairs; on quiet or flowing fresh or marine water; Halobates occurs on open oceans hundreds of miles from land; predatory generally on insects; oceanic forms feed on plankton and floating jellyfishes; about 350 species; in all zoogeographic regions and intervening oceans between 40° north and 40° south latitudes.Family Veliidae (smaller water striders)Habits and structures similar to Gerridae, but generally smaller, stouter bodied, and with hind femurs not reaching beyond tip of abdomen; two genera with last segment of middle tarsus split and with collapsible, fan-shaped tuft of long hairs used for propulsion over water surface; about 300 species; in all zoographic regions, genus Trochopus marine, but not in open sea.Family Hydrometridae (marsh treaders)Body, head, and legs long and slender; claws arise at tip of tarsus; wings present or absent; crawl weakly and clumsily among emergent plants in fresh water; predatory; more than 100 species; mostly tropical, a few species in North America and Eurasia.Family Mesoveliidae (water treaders)Small; body and legs slender; live on fresh water, especially among emergent plants or on saturated mud; one New Guinea species lives on the forest floor away from water; about 30 species; in all zoogeographical regions.Family Saldidae (shore bugs)Head large; eyes prominent; labium apparently 3-segmented, long, first segment very short and stout, second very long; legs long and slender for running or leaping; generally inhabit land surfaces near fresh or salt water; some species live on rocks in the intertidal zone, hide in air trapped in crevices when rocks are covered at high tide; predatory on small or crippled insects; over 200 species, in all zoogeographic regions.Family LeptopodidaeLike Saldidae; all 24 species native to Old World, one Mediterranean species established in New World (California).Family Hebridae (velvet water bugs)Tiny, covered with hydrofuge hairs suitable to habitat on surface or along edges of bodies of still, fresh water; sometimes crawl into the water; food habits not known; about 120 species; in all zoogeographic regions; some widely distributed species may have been carried in drinking water on early sailing ships.Family LeotichiidaeStructure suggestive of Saldidae but distinguished by strong median ridge on pronotum; habits unlike Hebridae; one of the two known species (both from southeastern Asia) occurs in caves.Suborder GeocorisaeTrichobothria either present on abdomen or absent from both abdomen and head; antennae longer than head; difficult suborder to define; perhaps a convenient ecological (rather than a taxonomic) category better replaced by elevation of the two series to suborders; terrestrial; habits and foods varied.Series CimicomorphaTrichobothria absent from head and abdomen; front wings with or without a cuneus (special region on hardened basal portion known as the corium).Family Miridae (plant bugs)Small, fragile; ocelli absent; labium 4-segmented; front wing with cuneus, membranous area with 1 or 2 closed cells; legs slender to stout, for walking, running, or jumping; predatory, phytophagous, or both; often host-specific; includes major crop pests (e.g., “lygus bugs” of the Northern Hemisphere, the “tea blight” [Helopeltis] in southeastern Asia), also several important predators of small insects; about 10,000 species; in all zoogeographic regions; found north of the Arctic Circle.Family Cimicidae (bedbugs)Oval to elongate oval, very flat; eyes small; forewings reduced to small, transversely oval pads exposing most of abdomen; hindwings absent; bloodsucking parasites of mammals and birds, usually present on host only while feeding, otherwise hiding in crevices in nest or lair; eggs glued to walls of crevices; contains the two species of bedbugs that attack man, Cimex lectularius, (originally in Europe and North America, now in all parts of the world) and Cimex hemipterus (common in southern Asia and Africa); C. lectularius probably encountered cave-dwelling men; not yet conclusively proved as carriers of any human diseases even though the bug can be successfully infected with some human parasites; about 75 species; found throughout the world but show marked parallels with ranges of their hosts.Family Reduviidae (assassin bugs)Head usually protruding; beak short, 3-segmented, strongly curved away from underside of head, with tip in deep stridular groove between front coxae; front legs adapted for grasping; prey on insects and other small animals; genus Triatoma and some close relatives suck blood of mammals (including man) and are carriers of the tropical American Chagas’ disease; can inflict painful bites; nearly 4,000 species; on all significant land masses.Family Anthocoridae (anthocorid bugs)Ocelli and cuneus present; membrane of wing with or without veins; male external genitalia asymmetrical; predatory, mostly on small arthropods, including their eggs; some eat ants; others inhabit bird nests; may bite man; about 400 species; in all zoogeographic regions.Family Polyctenidae (bat bugs)Oval; flattened; eyeless; front forewings, rudimentary, unhinged; hindwings absent; head and certain other parts with transverse rows of spines forming ctenidia (combs); eggs hatch within female, young nymphs retained in body and nourished by mother through a pseudoplacenta; bloodsuckers live entire life in fur of certain bats; 20 species; found from southern North America to South America, in Africa, and from southern Asia to Java.Family Aradidae (flat bugs)Extremely flat; surface usually rough; head horizontal; clypeus prominently convex; ocelli absent; legs short; tarsi two-segmented; abdomen much wider than wings when present; found in crevices, as under loose bark or debris on the ground; feed on molds or fungi; nearly 1,000 species in all zoogeographic regions; sometimes part of group separated as family Dysodiidae.Family Tingidae (lace bugs)Head small, without ocelli; antennae slender to stout, generally with first 2 segments very short, 3rd long; pronotum and forewings often finely to coarsely reticulate (lacy); tarsi 2-segmented; phytophagous, on anything from mosses to flowering plants, often host-specific; several species are common pests on maize, cotton, cacao, and other cultivated plants; almost 800 species; in all zoogeographic regions.Family Nabidae (nabid bugs)Ocelli present; beak 4-segmented, long, generally curved away from underside of head; no striated groove between front coxae of legs; front legs more or less thickened for grasping; forewings without a cuneus; predatory on soft-bodied insects; frequently investigated as a possible biological control agent for insect pests of field crops; about 300 species; in all regions of the world.Family Phymatidae (ambush bugs)Stout; surface roughened by tubercles; beak short, 3-segmented, curved away from underside of head, tip resting in stridular groove between front coxae and legs; 4th antennal segment conspicuously thickened; front femurs of legs strongly thickened; scutellum small and triangular or large, broad, and reaching tip of abdomen; predatory, generally, lying among flower petals awaiting prey; more than 200 species; found on all major continents except Australia; not found on oceanic islands.Family Enicocephalidae (unique-headed bugs)Predatory; about 130 species; in all zoogeographic regions.Family Dipsocoridae (or Cryptostemmatidae or Ceratocombidae)Biology poorly known; about 200 species; in all zoogeographic regions, more than half of species found in tropical America.Family IsometopidaePhytophagous; less than 60 species; sometimes reduced to a subfamily in the Miridae.Family JoppeicidaeThe sole species occurs in the eastern Mediterranean region.Family MicrophysidaePredatory; about 25 species; found in Eurasia and North America; 1 African genus questionably placed here.Family SchizopteridaeAbout 100 species; sometimes united with the Dipsocoridae (see above).Family TermitaphididaeLives with termites; includes 9 species; found in tropical America, Africa, India, and Australia.Family Thaumastocoridae (or Thaumastotheriidae)Phytophagous; about 12 species; found in Australia, India, North and South America, and the Greater Antilles.Family VelocipedidaePredatory; 4 species; in the Orient.Family VianaididaeLives with ants (myrmecophilous); 4 species; found in tropical America.Series Pentatomomorpha (or Trichophora)Trichobothria on abdomen; cuneus always absent; terrestrial; various habits and foods.Family Pentatomidae (stinkbugs)Head with side margins generally expanded over bases of antennae; ocelli usually present; antennae usually 5-segmented; scutellum large to very large, triangular or U-shaped; without claval commissure; tibiae without rows of spines; phytophagous or predatory, sometimes within same species; several important pests, including the southern green stinkbug (Nezara viridula), a serious pest of fruit and vegetable crops throughout the world; the harlequin cabbage bug (Murgantia histrionica), on North American cruciferous crops; the senn bugs (Eurygaster species), on small grains in the Old World—several genera (Podisus, Zicrona) feed on caterpillars and other larvae; about 5,000 species; in all zoogeographic regions; taxonomy unsettled, especially assignment of taxonomic levels; definition and relationships of subgroups not in dispute.Family PlataspidaeOval to transversely oval; head small, inserted deeply into prothorax; scutellum large, broadly U-shaped, reaching apex of abdomen; forewings much longer than body; wing membrane transversely folded at base and inserted under scutellum; tibiae not spined; phytophagous; about 500 species; found only in the Old World.Family Cydnidae (burrower bugs)Scutellum triangular or broadly U-shaped; forewings without claval commissure (except genus Amnestus); tibiae with rows of spines; colour generally brown or black, sometimes with creamy markings; phytophagous, most species in soil feed on roots of host; more than 750 species; in all zoogeographic regions; subfamily Thyreocorinae (or Corimelaeninae) sometimes given family status.Family CoreidaeAntennae 4-segmented, inserted on or above imaginary line connecting middle of eye to front end of buccula (cheek plates); ocelli present; forewings with claval commissure; wing membrane with numerous interconnected veins; appendages often with leaflike expansions; phytophagous, some as pests of cultivated plants, for example, the squash bug (Anasa tristis) on cucurbit crops in North America, a rice bug (Leptocorisa varicornis) in the Orient; more than 2,000 species; in all zoogeographic regions; subfamilies Alydinae and Rhopalinae (or Corizinae) sometimes given family status.Family Berytidae, or Neididae (stilt bugs)Body slender, narrowly elongate; legs extremely long, linear; labium 4-segmented; usually phytophagous, sometimes attack soft-bodied plant-feeding insects; more than 100 species; in all zoogeographic regions.Family PiesmatidaeJuga (2 outer lobes of head) extend as a pair of horizontal, hornlike projections; prothorax with a deep pit below expanded sides; forewings, except apical part of membrane, with numerous coarse punctures; phytophagous, most frequently on plants of the goosefoot family (Chenopodiaceae); proven vectors of viral diseases of sugar beets in Europe and North America; about 20 species; in all zoogeographic regions.Family LygaeidaeAntennae 4-segmented, inserted on side of head below an imaginary line connecting middle of eye to front end of buccula; ocelli present except in brachypterous forms; forewings with a claval commissure; wing membrane with 4 or 5 simple, longitudinal veins; phytophagous (on sap or seeds) or predatory (on mites and small insects); pests include the chinch bug (Blissus leucopterus) of North America, the Old World cotton stainer (Oxycarenus hyalinipennis), and several species of false chinch bugs in the widely distributed genus Nysius; in North America Geocoris, considered beneficial, feed upon mites and small insects attacking crop plants; contains between 3,000 and 5,000 species found in all parts of the world, including many small islands.Family LargidaeHead triangular, without ocelli; forewings with claval commissure; female with 6th visible abdominal segment below cleft medially; phytophagous; habits not well known; of no economic importance; about 100 species; in all zoogeographic regions.Family Pyrrhocoridae (fire bugs, cotton stainers)Head triangular, without ocelli; forewings with long claval commissure; female with 6th visible abdominal segment below entire; phytophagous or predatory; cotton stainers (Dysdercus†), serious pests of cotton crops; some species (e.g., genus Dindymus) feed on flies as adults, nymphs prefer termites; more than 300 species; in all zoogeographic regions.Family AphylidaeTwo species in Australia.Family ColobathristidaePhytophagous; about 90 species; inhabit Neotropical, Oriental, and Australian regions; unknown in North America, Europe, and Africa.Family HyocephalidaeOne species in Australia.Family LestoniidaeTwo species in Australia.Family PhloeidaeThree species in tropical America, 1 in Borneo.Family StenocephalidaeAbout 40 species; except for 1 species on the Galapagos Islands, all known species live in Eurasia and Africa.Family UrostylidaeApproximately 50 species confined to Asia and Australia.

Critical appraisal

The classification of the Heteroptera is in a state of flux due to continuous investigations in ontogeny, genetics, ecology, and morphology. Not all results are in agreement, suggesting that important changes probably will occur. Wide disparity in superfamily concepts has resulted in dropping them from the above classification of the heteropterans.

The division of the order into three suborders is generally accepted, even if only for convenience; the prefixes reflect the broad ecological habits of the families included. Hydrocorisae reflects the adaptation of most of its families (Gelastocoridae and Ochteridae excepted) for life under water. Amphibicorisae suggests the land and water habits of these insects that live on the surface of the water or along shores. Geocorisae reflects the preferred terrestrial habitats of members of this group.